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Microsaccades are an important component of the small eye movements that constitute fixation, the basis of visual perception. The specific function of microsaccades has been a long-standing research problem. Only recently, conclusive evidence emerged, showing that microsaccades aid both visual perception and oculomotor control. The main goal of this thesis was to improve our understanding of the implementation of microsaccade generation within the circuitry of saccade control, an unsolved issue in oculomotor research. We make a case for a model according to which microsaccades and saccades result from mutually dependent motor plans, competing for expression. The model consists of an activation field, coding for fixation at its center and for saccades at peripheral locations; saccade amplitude increases with eccentricity. Activity during fixation spreads to slightly peripheral locations in the field and, thus, may result in the generation of microsaccades. Inhibition of remote and excitation of neighbouring locations govern the dynamics of the field, resulting in a strong competition between fixation and saccade generation. We propose that this common-field model of microsaccade and saccade generation finds a neurophysiological counterpart in the motor map of the superior colliculus (SC), a key brainstem structure involved in the generation of saccades. In a series of five behavioral experiments, we tested implications of the model. Predictions were derived concerning (1) the behavior of microsaccades in a given task (microsaccade rate, amplitude, and direction), (2) the interactions of microsaccades and subsequent saccades, and (3) the relationship between microsaccadic behavior and neurophysiological processes at the level of the SC. The results yielded strong support for the model at all three levels of analysis, suggesting that microsaccade statistics are indicative of the state of the fixation-related part of the SC motor map.
We present an approach to generate (multivariate) twin surrogates (TS) based on recurrence properties. This technique generates surrogates which correspond to an independent copy of the underlying system, i.e. they induce a trajectory of the underlying system starting at different initial conditions. We show that these surrogates are well suited to test for complex synchronisation and exemplify this for the paradigmatic system of Rossler oscillators. The proposed test enables to assess the statistical relevance of a synchronisation analysis from passive experiments which are typical in natural systems
When the eyes fixate at a point in a visual scene, small saccades rapidly shift the image on the retina. The effect of these microsaccades on the latency of subsequent large-scale saccades may be twofold. First, microsaccades are associated with an enhancement of visual perception. Their occurrence during saccade target perception could, thus, decrease saccade latencies. Second, microsaccades are likely to indicate activity in fixation-related oculomotor neurons. These represent competitors to saccade-related cells in the interplay of gaze holding and shifting. Consequently, an increase in saccade latencies would be expected after microsaccades. Here, we present evidence for both aspects of microsaccadic impact on saccade latency. In a delayed response task, participants made saccades to visible or memorized targets. First, microsaccade occurrence up to 50 ms before target disappearance correlated with 18 ms (or 8%) faster saccades to memorized targets. Second, if microsaccades occurred shortly (i.e., < 150 ms) before a saccade was required, mean saccadic reaction time in visual and memory trials was increased by about 40 ms (or 16%). Hence, microsaccades can have opposite consequences for saccade latencies, pointing at a differential role of these fixational eye movements in the preparation of saccade motor programs
Fixational eye movements occur involuntarily during visual fixation of stationary scenes. The fastest components of these miniature eye movements are microsaccades, which can be observed about once per second. Recent studies demonstrated that microsaccades are linked to covert shifts of visual attention. Here, we generalized this finding in two ways. First, we used peripheral cues, rather than the centrally presented cues of earlier studies. Second, we spatially cued attention in vision and audition to visual and auditory targets. An analysis of microsaccade responses revealed an equivalent impact of visual and auditory cues on microsaccade-rate signature (i.e. an initial inhibition followed by an overshoot and a final return to the pre-cue baseline rate). With visual cues or visual targets, microsaccades were briefly aligned with cue direction and then opposite to cue direction during the overshoot epoch, probably as a result of an inhibition of an automatic saccade to the peripheral cue. With left auditory cues and auditory targets microsaccades oriented in cue direction. We argue that microsaccades can be used to study crossmodal integration of sensory information and to map the time course of saccade preparation during covert shifts of visual and auditory attention